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23 November 2018 Taxonomic Significance of Petal and Sepal Accepted author version posted online: 23 November 2018 Taxonomic significance of petal and sepal micromorphological characteristics in some Justicia Nees (Acanthaceae) species in Peninsular Malaysia 1,2Amirul-Aiman, A.J., 1Noraini, T., 3Nurul-Aini, C.A.C., 1Khaulah, A.R., 1,4Nurul-Syahirah, M., 1,5Amalia, R. and 1Ruzi, A.R. 1School of Environmental Science and Natural Resources, Faculty of Science and Technology, Universiti Kebangsaan Malaysia, 43600 UKM Bangi, Selangor, Malaysia 2Department of Environmental Sciences, Faculty of Environmental Studies, Universiti Putra Malaysia, 43400 UPM Serdang, Selangor, Malaysia 3Department of Plant Science, Kuliyyah of Science, International Islamic University Malaysia, Bandar Indera Mahkota, 25200 Kuantan, Pahang, Malaysia 4Department of Agronomy, Faculty of Plantation and Agrotechnology, Universiti Teknologi MARA, Jasin Campus, 77300 Merlimau, Melaka 5Department of Biology, Mathematics and Science Faculty, Syiah Kuala University, 23111 Darussalam, Banda Aceh, Indonesia *Corresponding author: [email protected] Running title: Petal and sepal micromorphological characteristics Abstrak: Acanthaceae merupakan famili tumbuhan berbunga di bawah order Lamiales dan terdiri daripada sekurang-kurangnya 4000 spesies di kawasan tropika dan juga kawasan subtropika dengan genus Justicia merupakan genus terbesar. Kajian mikromorfologi petal dan sepal telah dijalankan ke atas ke atas enam species iaitu J. betonica L., Justicia carnea Lindl., J. comata (L.) Lam., J. gendarussa Burm.f., J. procumbens L. and J. ptychostoma Nees. Kaedah kajian termasuklah penetapan bunga di dalam larutan penetap, siri dehidrasi, titik pengeringan genting dan cerapan di bawah mikroskop imbasan elektron. Hasil kajian menunjukkan kesemua enam spesies mempunyai variasi dan perbezaan mikromorfologi pada permukaan epidermis petal dan sepal. Kutikel lilin boleh ditemui pada kedua-dua permukaan epidermis petal dan sepal kesemua spesies yang dikaji. Kutikel lilin butir merupakan satu-satunya jenis kutikel lilin yang ditemui pada kesemua permukaan petal dan sepal spesies kajian. Terdapat lapan jenis trikom ditemui pada permukaan epidermis sepal dan juga lapan jenis trikom pada permukaan epidermis petal. Justicia betonica boleh dibezakan secara langsung dengan kehadiran stomata amfidiasitik pada permukaan epidermis petal dan stomata tidak hadir pada permukaan petal spesies yang lain. Kesimpulannya, hasil kajian ini membuktikan ciri mikromorfologi petal dan sepal mempunyai kesignifikan sistematik dan boleh digunakan sebagai data sokongan bagi pembezaan dan pengenalpastian spesies dalam genus Justicia. Kata kunci: mikromorfologi sepal dan petal, Justicia, Acanthaceae Abstract: Acanthaceae is a flowering plants family under the order Lamiales and comprises at least 4000 species regardless of tropical and subtropical species with Justicia being the largest genus under this family. The study on the flower’s petal and sepal micromorphology have been done on six Justicia species, namely as J. betonica L., Justicia carnea Lindl., J. comata (L.) Lam., J. gendarussa Burm.f., J. procumbens L. and J. ptychostoma Nees. Methodology used involves fixation, dehydration series, critical point drying technique and observation under scanning electron microscope. Results have shown that all six species have differences in micromorphological characteristics in the sepals and petals epidermal surfaces. Cuticular wax can be found on both abaxial and adaxial epidermal surfaces of petal and sepal for all species studied. Granular cuticular wax is the only type of cuticular wax was found present on petal and sepal of all species studied. There are also eight types of trichomes present on the petal epidermal surface and eight types of trichomes present on the sepal epidermal surface. J. betonica can be recognized directly by having amphidiacytic stomata on petal epidermal surface while stomata were absent on the petal epidermal surface of other species. As a conclusion, findings in this study have proven that petal and sepal micromorphological characteristics have systematic significant and can be used as supporting data for species differentiation and identification in the genus Justicia. Keywords: sepal and petal micromorphology, Justicia, Acanthaceae INTRODUCTION Acanthaceae is a flowering plant family under the order of Lamiales and comprises at least 4000 species regardless of tropical and subtropical species (Borg 2008). According to Hu et al. (2011) there are about 220 genera and 4000 species in the pantropical and subtropical region with few species can be found in the temperate region. About 35 genera are native in Peninsular Malaysia and few have been brought to Peninsular Malaysia as ornamental plants. Acanthaceae is also the third largest tropical plants after the Myrtaceae and Melastomataceae (Grant 1995). Acanthaceae comprises of three subfamilies which are Acanthoideae, Thunbergioideae and Nelsonioideae (Melchior 1964; Keng 1986; Borg & Schonenberger 2011). Then, Scotland and Vollesen (2000) classified Acanthaceae into three subfamilies, Nelsonioideae, Acanthoideae and Thunbergioideae. While McDade et al. (2008) consider the family consists of four subfamilies, Acanthoideae, Nelsonioideae, Thunbergioideae and Andrographideae. But later, Vollesen (2008) elevated the Tribe Ruellieae to subfamily Ruellioideae. New findings by Schwarzbach and McDade (2002) and Borg (2008) have suggested that Avicennia has a sister relationship with Acanthaceae but still questionable and unclear in Acanthaceae lineages. The species from the Acanthaceae can be found in many habitat, having lots of morphological characters and biogeographical distribution. Taxonomic studies of Asian Justicia have been done by few researchers such as Clarke (1885), Benoist (1935), Hansen (1989), Cramer (1998) and Ruengsawang et al. (2012), but the taxonomical study on the Justicia in Malaysia is still scarce with only researches by some researchers such as Ridley (1923) and Ruth (2009) been known. In 2017, a study on current morphology-based classification of “justicioid” lineage with focus on Old World taxa by Kiel et al. (2017) confirmed a strong support of monophyly of New World justicioids including all New World species of Justicia. They also added that even though some genera of justicioids are monophyletic, Justicia is grossly polyphyletic. The results were obtained by the combinations of diagnostic utility of morphological characters for major clades focusing on the inflorescences, androecia, seeds and pollen. Justicia is the largest genus in Acanthaceae with about 600 species and distributed in the tropical and subtropical as well in both hemisphere (Wasshaushen 1989). Mabberley (2008) stated that, Justicia L. is one of the largest and most complex genera in Acanthaceae. According to McDade et al. (2008), even though Acanthaceae is an important component in the tropical and subtropical all over the world, taxonomists are still having scarce information about the family especially in part of petal and sepal micromorphology. In this study, the investigation focused just at the micromorphological characteristics of petal and sepal epidermal surfaces on some Justicia species in Peninsular Malaysia in order to understand more on the micromorphological characteristics of its’ flower (especially on the petals and sepals). MATERIALS AND METHODS Six species were chosen (J. betonica T. Anderson, J. carnea Hook. ex Nees, J. comata Vellozo ex Nees, J. gendarussa J. Macrae ex Nees, J. procumbens T. Anderson ex Nees, and J. ptychostoma Nees). Fresh samples of flower were used in this study and were collected from several forests reserved as shown in Table 1 and Figure 1. Observation under scanning electron microscope follows Barthlott et al. (1998) and Noraini (2006) with few modifications. Small piece of petal and sepal were fixed in 4% glutaraldehyde for 12-24 hours at 4oC prior to the dehydration in alcohol series (35%, 50%, 75%, 95%, 100%, 100% % 100%) followed by critical point drying using drying machine model Polaron CPD 7501, were then been attached to the aluminium stubs with double sided adhesive tape and then sputter-coated with a thin film of gold using Gold Coater model BioRad SC500 to improve the electrical conducting properties of the leaf surface. Samples then were examined under a scanning electron microscope model Philips XL Series XL 30 with magnification up to 10000x. The description of petal and sepal micromorphology follows Radford et al. (1974) and Singh and Jain (1975). RESULTS Petal Justicia comata Wax: present in form of film layer and granular on both abaxial and adaxial epidermal. Cuticular striation on abaxial epidermal: obscure, periclinal and anticlinal wall cannot be differentiated. Cuticular striation on adaxial epidermal: clearly define, with periclinal wall mergedn at certain area, forming conical shape papillae, anticlinal wall sunken, tiny striae present at certain area of epidermal surface and present almost parallel towards the peak of papillae, no striae present at the peak of papillae. Stomata: absent. Trichome: simple multicellular or simple uniseriate (short to long, with echinate ornamentation) sparsely scattered on the abaxial epidermal and peltate glandular trichome (multicellular terminal – 2 cells) sparsely scattered on the adaxial epidermal. Justicia betonica Wax: present in form of film layer and flakes cuticular waxes on the abaxial epidermal, film and granular cuticular waxes present on the adaxial epidermal. Cuticular striation
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